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The International Journal of Plant Reproductive Biology 12(1) Jan., 2020, pp.44-55

DOI 10.14787/ijprb.2020 12.1.

Karyological and Cytogenetical Studies on Gymnosperms in V.N. Sukachev Institute of Forest

Elena N. Muratova*, Tamara S. Sedel’nikova, Olga V. Goryachkina and Alexander V. Pimenov

V.N. Sukachev Institute of Forest, Russian Academy of Sciences, Siberian Branch, Federal Research Center «Krasnoyarsk Science Center SB RAS», Akademgorodok 50/28, Krasnoyarsk, 660036, Russia

Universitetskaya nab. 7/9, Russia.

*Corresponding author e-mail: [email protected];

Received :

ABSTRACT

Keywords :

02.12.2019; Accepted and published online: 15.12.209

More than 200 populations and provenances of representatives of different genera from the families Pinaceae and Cupressaceae, and also from the genus Ephedra, family Ephedraceae, were studied. Investigations were carried out in natural populations and at introduction, in different environmental conditions, in botanical gardens and parks; various intraspecific forms and unique trees have been studied. The variability of chromosome numbers and a wide range of chromosomal mutations have been revealed. The studies on karyotypes in species of conifers showed higher level of chromosomal anomalies and their wide spectrum in extreme conditions, at introduction, in cultivars and abnormal form of trees. Use of fluorescence in situ hybridization (FISH) with the 45S and 5S ribosomal RNA gene probes and DAPI staining allows to identify of homologous chromosome pairs in the karyotypes of conifers and to facilitate the comparative karyotype analysis of these species.

populations, intraspecies forms, introduction, conifers, karyotype, chromosomes, variability, chromosomal mutations, ribosomal RNA genes.

EPR R OT DN UAL CP T IF VEO BYT IOEI L

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Karyological and cytogenetic studies on Gymnosperms others). Fluorescence in situ hybridization (FISH) was done make a great contribution to the knowledge of their biotic according to a previously published protocol (Badaeva et al. diversity, to the solution of systematics, evolutionary and 1996, Goryachkina et al. 2013a).population genetics problems. At V.N. Sukachev Institute of

RESULTS AND DISCUSSIONForest SB RAS researches of conifer karyotypes started by

Studies cover species from 4 genera of family Pinaceae L.F. Pravdin in the 60s of XX century with the study of Scots Spreng. ex F. Rudolphi: Abies Mill. (True fir), Larix Mill. pine Pinus sylvestris L. (Pravdin 1964). In the late 60s – early (Larch), Picea A. Dietr. (Spruce), Pinus L. (Pine); 6 genera of 70s karyological research at the Institute was carried out by the family Cupressaceae Bartl.: Chamaecyparis Spach (M.V. Kruklis (Kruklis 1971a, b, 1974a, b, Kruklis and

), Cupressus L. ( ), Juniperus L. (Juniper), Milyutin 1977). She studied the karyotypes of several larch Microbiota Kom., Sequoiadendron Buchh., Thuja L.; the species from Siberia – Larix sibirica Ledeb., L. gmelinii genus Ephedra L., family Ephedraceae Dumort. Total more (Rupr.) Rupr. and L. czekanowskii Szaf. (L. sibirica × L. than 200 populations and provenances were studied. Investi-gmelinii), meiosis in these species, karyotype of Siberian gations were carried out in natural populations and at intro-spruce (Picea obovata Ledeb.). Furthermore, M.V. Kruklis duction, in different environmental conditions (optimum and was one of the first authors who found additional extreme, disturbed ecosystems), in botanical gardens and chromosomes (B-s) in conifers (Kruklis 1971a, b). Since the parks; various intraspecific forms and unique trees have been 70s and until now karyological studies of different species of studied.conifers growing in various climatic regions, mainly in the

Karyotypes of all the examined Abies species from boreal forests of Eurasia are in progress.Russia and neighboring countries [A. sibirica Ledeb., A. alba

MATERIALS AND METHODSMill., A. holophylla Maxim., A. lasiocarpa (Hook.) Nutt., A. sachalinensis (F. Schmidt) Mast., A. semenovii B. Fedtsch., A. Materials for investigations were seeds of different veitchii Lindl.] include 24 chromosomes (2n = 24). Diploid gymnosperm species collected in natural populations and at complement of True fir consists of 7 pairs of long symmetric introduction, in optimal and extreme conditions, in disturbed (metacentric) and 4 pairs of asymmetric (submeta- and ecosystems, botanical gardens and parks, in various subacrocentric) chromosomes (Muratova et al. 1991, 1998, intraspecific forms and unique trees. Classic and new methods 2001, 2008, Muratova and Matveeva 1996, Muratova 1997a, for karyological and cytogenetical studies were used. Pimenov and Sedel’nikova 2002, 2003, Sedel’nikova and Pretreatment of the germinated seeds, fixation, staining of Pimenov 2003, 2005a, Kvitko and Muratova 2009, 2010, slides with acetohematoxylin was performed according to Kvitko et al. 2009, 2011, Sedel’nikova et al. 2011, standard for coniferous methods (Pravdin et al. 1972, Goryachkina et al. 2013b, Sedel’nikova 2014). Muratova 1978, 1979a, b, 1995a, Muratova et al. 2008 and

False cypress Ñypress

Studied Abies species differ on number and localization Meiosis during microsporogenesis in Siberian fir in of secondary constrictions on chromosomes. Chromosome natural stands and at the arboretum of the V. N. Sukachev complement of Siberian fir (A. sibirica) is given on Fig.1. Institute of Forest has been studied in details (Bazhina et al. Karyological studies on Siberian fir (A. sibirica) populations 2006, 2007a-c, 2008, 2011). Features of A. sibirica meiosis from different parts of area revealed a relatively low level of were revealed. Different stages of meiosis are given on Fig. 2. intraspecific karyological polymorphism and slight Meiotic irregularities were found at different stages. Common differences in features of nucleolar regions localization and specific types of irregularities have been described in (Muratova and Matveeva 1996, Sedel’nikova and Pimenov studied trees. The range of irregularities under the arboretum 2005a, Kvitko and Muratova 2009, 2010, Kvitko et al. 2009, conditions was much wider than in natural ecosystems. It is 2011). proposed that some meiotic irregularities can be cause of

pollen sterility. Investigated Larix species include L. decidua Mill., L.

sibirica, L. gmelinii, L. ×czekanowskii (hybrid complex between L. sibirica and L. gmelinii), L. sukaczewii Dylis, L. cajanderi Mayr, L. ×ochotensis Kolesn. (possibly hybrid between L. cajanderi and L. kurilensis Mayr) and L. ×amurensis Kolesn. (supposedly L. gmelinii, L. maritima Suk. and L. olgensis A. Henry). Chromosome numbers in the seed progeny of typical habitus L. sibirica trees, intraspecific forms, differing in the color of female cones, as well as morphotypes deviating from normal ones – with “witch brooms”, bush-like, with large and small cones were determined (Muratova and Chubukina 1985, Muratova 1991a, b, 1993, 1994, Muratova et al. 2007, 2008, Pimenov and Sedel’nikova 2002, 2003,

Figure 1. Karyotype of Abies sibirica Ledeb., 2n = 24. The Sedel’nikova and Pimenov, 2005b, 2007, 2017a, b, 2019, chromosomes are distributed into groups according to results of the Sedel’nikova et al. 2005, 2011; Syzikh et al. 2006; morphometric karyotype analysis. I-XII indicates the numbers of the

Vladimirova et al. 2007, Kvitko et al. 2009, Sedel’nikova chromosomes. Material stained with acetohematoxylin. Scale bar – 10 µm. 2014).

arose as a result of crossing

Fig. 2. Meiosis and pollen formation in Siberian fir Abies sibirica Ledeb.: 1 – leptotene, 2 – pachytene, 3 – diakinesis, 4 – metaphase I, 5 – anaphase I, 6 – telophase I, 7 – prophase II, 8 – metaphase II, 9 – telophase II, 10 – tetrad of nuclei, 11 – tetrad of microspores, 12 – pollen grains. Magnification: Material stained with acetohematoxylin. 10x60.

1

7 8 9 10 11 12

2 3 4 5 6

2020 45Karyological and Cytogenetical Studies on Gymnosperms in V.N. Sukachev Institute of Forest

Diploid complement of Larix species includes 24 morphology of some Picea species showed different types of chromosomes (2n = 24). Their karyotypes contain six pairs of irregularities in processes of meiosis and pollen development metacentric and six pairs of submeta- or intercentric (Bazhina et al. 2017, 2019).

Detailed karyological and cytogenetical researches of chromosomes (Kruklis 1974a, Muratova and Chubukina Siberian spruce from different population of Siberia were 1985, Muratova 1991b, 1993, 1997a, Muratova et al. 1991, carried out (Kruklis 1971a, b, Medvedeva and Muratova 1987, 2007, 2008, Sedel’nikova and Pimenov 2005b, 2019, Syzikh Broka 1990, Muratova 1997a, Muratova and Vladimirova et al. 2006, Goryachkina et al. 2013a). Chromosome 2001b, Vladimirova 2002, Sedelnikova et al. 2004, 2011, complements of L. sibirica and L. gmelinii are given on Fig. 3. Vladimirova and Muratova, 2005, 2006, Muratova et al. 2008, Borisov and Muratova 2010). B-chromosomes (2n = 24+1-4B) in many populations and provenances of this species have been revealed.

In addition to Siberian spruce, one or more B-chromosomes include other Picea species studied by us from Russia, Belarus, Kazakhstan, Canada, Kyrgyzstan, China, USA, France, Czech Republic, Japan. They are: P. abies (L.) Karst. – 1-4B, P. ajanensis (Lindl. et Gord.) Fisch. ex Carr. – 1-3B, P. breweriana S. Wats. – 1Â, P. ×fennica (Regel) Kom. (hybrid between P. abies and P. obovata) – 1Â, P. glauca

Fig. 3. Chromosome complements of Larix sibirica Ledeb. (a) and L. (Moench.) Voss – 1-3B, P. glehnii (Fr. Schmidt) Mast. – 1-5B, gmelinii (Rupr.) Rupr. (b), 2n = 24. Magnification: Material P. koyamae Shiras. – 1Â, P. meyeri Rehder & E.H. Wilson – 1-stained with acetohematoxylin.

3B, P. pungens Engelm. – 1Â, P. purpurea Mast. – 1Â, P. Detailed karyological studies of different populations of sitchensis (Bong.) Carr. – 1-2B, P. schrenkiana Fisch. et C. A.

widespread boreal species L. sibirica, L. sukaczewii, L. Mey. – 1B (Muratova and Frolov 1995, Muratova and gmelinii, L. cajanderi were carried out. It was established that Sedel’nikova 2000, Muratova and Vladimirova 2001a, different Larix species and populations differ in the number Vladimirova et al. 2003, 2007, Muratova et al. 2004, and localization of secondary constrictions. For the first time Sedel’nikova et al. 2011, Karpyuk and Muratova 2005, among representatives of this genus, we discovered Vladimirova et al. 2007, Kvitko et al. 2009, Pimenov et al. additional, or B-chromosomes (2n = 24+1B) in L. gmelinii in 2012, Goryachkina et al. 2013b, Tashev et al. 2014, 2015, Eastern Siberia and L. sibirica in Soutern Siberia and on the Muratova 2018). To date, 21 species with B-chromosomes, north of Middle Siberia (Muratova 1991a, 1994, Sedel’nikova including interspecific hybrid P. ×fennica, have been found in et al. 2005, 2011, Syzikh et al. 2006, Muratova et al. 2007, the genus Picea. Scientists of V.N. Sukachev Istitute of Foresr 2008, Sedel’nikova and Pimenov, 2007, Kvitko et al. 2009). for the first time found B-chromosomes in P. abies, P.

Studies on meiosis in L. gmelinii at the V.N. Sukachev breweriana, P. obovata, P. pungens, P. purpurea, P. Institute of Forest Arboretum showed that this process was schrenkiana. Chromosome complements of some Picea characterized by various anomalies. Common and specific species are given on Fig. 4. types of meiotic irregularities were found at different stages. The studied species of the genus Pinus belong to two These irregularities can be the cause of heterogeneous pollen subgenera: Strobus (section Strobi, subsection Cembrae) and formation with different ploidy levels and other anomalies of Pinus (section Pinus, subsection Sylvestres). In Cembrae development (Goryachkina and Muratova 2016). group Siberian stone pine P. sibirica Du Tour, Korean stone

Karyotypes of studied Picea species include 24 pine P. koraiensis Siebold et Zucc., Siberian dwarf pine P. chromosomes (2n = 24): eight pairs of the long metacentric, pumila (Pall.) Regel from Russia and European stone pine P. two pairs of the short metacentric and two pairs of short cembra L. from Ukraine and Czech Republic were analyzed submetacentric chromosomes (Kruklis 1971a, b, Medvedeva (Muratova, 1978, 1979a, b, 1980, 1983, 1997a, Muratova et and Muratova 1987, Muratova et al. 1991, 1998, 2001, 2004, al. 1998, Sedel’nikova et al. 1999, Muratova and 2008, Muratova and Frolov 1995, Muratova 1997a, 2003, Sedel’nikova 2000, Sedel’nikova and Muratova 2002, Muratova and Vladimirova 2001a, b, Vladimirova 2002, Muratova et al. 2008, Sedel’nikova 2014). Vladimirova et al. 2003, 2007, Sedelnikova et al. 2004, 2011, In Sylvestres group P. sylvestris (Scots or Scotch) pine Karpyuk et al. 2005, 2009, Karpyuk and Muratova 2005, from Russia, Bulgaria and Czech Republic was studied in Kvitko et al. 2009, Sedel’nikova 2014, Goryachkina et al. details. In Scots pine, not only typical trees were studied, but 2018). Studies on meiosis during microsporogenesis in also individuals with deviating from normal habitus – dwarfs, Siberian spruce (P. obovata) and pollen development and with a weeping crown shape, with “witch brooms” and so on

10x90.

a b

46 The International Journal of Plant Reproductive Biology 12(1) Jan., 2020, pp.44-55

(Muratova 1983, Sedel’nikova and Muratova 1991, 1992, hybrid individuals. Mixoploidy was revealed in the seed 2001, Muratova and Sedel’nikova 1993, 2000, Sedel’nikova progeny of some species from the Arboretum Sofronka (near et al. 2000, 2011, Muratova et al. 2001, 2008, Pimenov and Plsen, Czech Republic): introduced from Serbia P. pinaster, Sedel’nikova 2002, 2003, Sedel’nikova 2003, 2014). Other from Spain P. uncinata Mill. ex Mirb., interspecific hybrid species of pine were analyzed also: P. pityusa Steven and P. between P. contorta Dougl. ex Loud. × P. banksiana Lamb.; in eldarica Medw. from Georgia, P. mugo Turra from Czech P. pinaster from park in Abkhazia (Sedel’nikova et al. 2008, Republic and Bulgaria, P. jeffreyi Grev. et Balf. from USA, P. Sedel’nikova 2016). Metaphase plates of some Pinus species pinaster Aiton from , P. heldreichii Christ. and P. are given on Fig. 5. nigra Arnold from Bulgaria (Muratova 1985, Pimenov and Sedel’nikova 2003, Sedel’nikova et al. 2005, 2008, 2011, Pimenov et al. 2012).

Karyotypes of studied Pinus species include 24 chromosomes (2n = 24). Pines of Cembrae group contain eleven pairs of long metacentric and one pair of short submetacentric chromosomes. Pines of Sylvestres group have ten pairs of long metacentric and two pairs of short submetacentric ones. Different Pinus species and populations differ in the number and localization of secondary constrictions (Muratova 1978, 1979a, b, 1980, 1983, 1997a, Sedel’nikova and Muratova 1991, 1992, 2001, 2002, Sedel’nikova et al. 1999, 2000, 2001, Muratova and Sedel’ nikova 1993, 2000, Muratova et al. 2008, Sedel’nikova 2014).

An increase in the variability of the chromosome numbers Fig. 5. Metaphase plates of Pinus jeffreyi Grev. et Balf., 2n = 24 (a) and in Pinus species, as well as other representatives of the family P. sylvestris L., 2n = 48 (b). Magnification: Material stained with

acetohematoxylin.Pinaceae, can occur during their introduction, as well as in

Abkhazia

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Fig. 4. Chromosome complements of Picea species: a – P. abies (L.) Karst., 2n = 24; b – part of P. abies metaphase plate with B -chromosome; c – P. koyamae Shiras., 2n = 24; d – P. koyamae, 2n = 24+1B; e – P. obovata Ledeb., 2n = 24; f – P. meyeri Rehder & E.H. Wilson, 2n = 24+3B. Arrows point to B-chromosomes. Scale bar 10 µm. Material stained with acetohematoxylin.

a

d

b

e

c

f

bf


Genome mutations, such as aneuploidy, mixoploidy, Special attention was paid to the karyological study of the Pinaceae family species on dry lands and swamps of various and, in some cases, polyploidy, were found in the studied types in the southern taiga subzone of Western Siberia species of Pinus, Larix, Picea, and Abies. Mixoploidy was (Sedel’nikova and Muratova 1991, 1992, 2001, 2002, found in L. sibirica and P. sylvestris tree forms abnormal in habit Muratova and Sedel’nikova 1993, Sedel’nikova et al. 1999, (Muratova and Sedel’nikova 2000, Sede’nikova et al. 2001, 2000, 2001, 2004, Sedel’nikova and Pimenov 2003, 2005a, b, 2011, Pimenov and Sedel’nikova 2002, 2003, Sedel’nikova Sedel’nikova 2014). Mixoploidy and aneuploidy were 2003, Muratova et al. 2007, 2008, Sedel’nikova and Pimenov detected in all studied species. The spectrum of genome and 2007, 2017a, b, 2019). In populations near the borders of areas, chromosome mutations in metaphases and ana-telophases in under extreme conditions, changes in the chromosome trees in the swamps was wider than in dry lands. Aberrations morphology, an increased occurrence of secondary were represented by ring and polycentric chromosomes, constrictions and chromosomal rearrangements are noted. fragments, irregularities of chromosome spirals. Multipolar A wide range of chromosome mutations was detected in mitoses, lagging chromosomes, chromosome bridges, Scots pine near the southern and northern borders of the range. fragments, ejections of chromosomes beyond the plate, Among them there are ring and polycentric chromosomes, agglutination of chromosomes were noted. deletions, fragments, multiple irregularities. The same trees

Among representatives of the family Cupressaceae a have anomalies during processes of mitosis and meiosis karyological study of Juniperus communis L. was carried out (Suntsov 1982a, b, Muratova 1991c, 1995b, 1997b, Muratova and the chromosome numbers of many species were and Sedel’nikova 2000, Sedel’nikova 2003). Chromosome determined. Cytological examination of Common juniper (J. mutations were found in Siberian larch (L. sibirica) in communis) on swamps and dry lands of Western Siberia Kazakhstan, Mongolia and in Taimyr under the influence of showed that the karyotype of this species contains 22 technogenic emissions, in Sukachev larch (L. sukaczewii) in chromosomes (2n = 22). All chromosomes were metacentric, the Southern Urals, in northern populations of Gmelin larch but one pair was close to the submetacentric type (Fig. 7). No (L. gmelinii), Siberian dwarf pine (P. pumila), Siberian spruce significant differences in karyotypes of swamp and dry valley (P. obovata), and Siberian fir (A. sibirica) in the mountains of populations were found (Mikheeva and Muratova 2005, Khamar Daban and high mountains of the Western Sayan Mikheeva et al. 2005, Muratova et al. 2008).(Suntsov 1982a, b, Muratova and Chubukina 1985, Muratova

and Matveeva 1996, Muratova 1991b, 1994, Muratova and Sedel’nikova 2000, Muratova et al. 2008, Kvitko and Muratova 2009, 2010, Sedel’nikova et al. 2011). Some examples of chromosome mutations in Pinaceae species (ring chromosomes and dicentric chromosome) are given on Fig. 6.

Fig. 7. Karyotype of Juniperus communis L., 2n = 22 (a); the chromosomes are distributed into groups according to results of the morphometric karyotype analysis (b). Material stained with acetohematoxylin.

Fig. 6. Chromosome mutations in Pinaceae species: ring chromo-Species widespread at introduction such as somes in Larix sibirica Ledeb. (a) and L. gmelinii (Rupr.) Rupr. (b);

Chamaecyparis lawsoniana (A. Murr.) Parl., Cupressus dicentric chromosome in P. koraiensis Nakai (c). Arrows show chromosome mutations. Magnification: Material stained with arizonica Greene, C. sempervirens L., Sequoiadendron acetohematoxylin.

10x90.

a b

c

a

b


giganteum (Lindl.) Buchh. (syn. Sequoia gigantea Torr.), cytogenetic markers makes it possible to identify Thuja îrientalis L., cultivars of T. occidentalis L. with chromosomes in the karyotype, select homologous pairs, and different color of needles and crown shape such as ‘Lutea’, identify chromosome mutations.

At present fluorescence in situ hybridization (FISH) with ‘Wareana’, ‘Wareana Lutescens’, ‘Globosa’ were studied (Pimenov and Sedel’nikova 2003, Sedel’nikova et al. 2005, 45S and 5S rRNA gene probes has been successfully used to 2008, 2011, 2014, Sedel’nikova, 2016). Seeds for researches study chromosomes of the main forest-forming coniferous were collected in parks and arboretums of Russia, Bulgaria, species in Siberia. Karyotypes of three Larix species (L. Kyrgyzstan, Ukraine, USA. The studied species have 22 sibirica, L. gmelinii, and L. cajanderi) were analyzed using chromosomes (2n = 22). These species are characterized by FISH and DAPI staining (Goryachkina et al. 2013a). Two mixoploidy (Fig. 8). major 45S ribosomal DNA loci per haploid genome have been

observed in the intercalary regions of two chromosomes (III

and IV) of L. sibirica, III, IV and VII chromosomes of L.

gmelinii and L. cajanderi. In addition to them, minor

nucleolus organizing regions (NORs) were mapped in

pericentromeric regions of chromosomes I, II, VI, and XII.

Two closely related species, L. gmelinii and L. cajanderi,

showed similar hybridization patterns. Only one locus of the

5S rDNA was found in all larch species in the distal region of

the chromosome III. This chromosome containing major loci

of the two ribosomal RNA gene families can serve as a marker

of the genus Larix (Hizume et al. 1995, Lubaretz et al. 1996,

Liu et al. 2006, 2007, Zhang et al. 2010, Goryachkina et al.

2013a). Karyotype analysis of Picea schrenkiana was carried out

using modern molecular cytogenetics methods (Goryachkina

et al. 2018). DAPI-bands were detected on 5 pairs of the

chromosomes. Major loci of 45S rRNA genes were revealed

on 5 pairs of the chromosomes; other 2 chromosome pairs with Fig. 8. Metaphase plates of Thuja occidentalis L. cultivars: 'Wareana',

2 minor sites of 45S rDNA were observed. Clusters of 5S 2n = 22 and 2n = 33 (a); 'Lutea', 2n = 22, 'Wareana Lutescens', 2n = rRNA genes occur on two arms of chromosome III; this 22. Magnification: Material stained with acetohematoxylin.

chromosome contains major locus of 45S rDNA near the M. decussata Kom. from Botanical Garden-Institute FEB

signal of 5S rDNA (Fig. 9). RAS (Vladivostok), a representative of the monotypic genus Microbiota, is also characterized by the number of chromosomes 2n = 22 (Kvitko et al. 2009). A karyological

It was established that they differ in the number of study of E. distachya L. from France (the genus Ephedra)

major loci of 45S rDNA in the intercalary regions of the of showed that this species is a tetraploid with 2n = 4x = 28

chromosomes: in P. obovata there are 6 of them, in P. (Muratova et al. 2001).schrenkiana 5 (Fig. 10). In P. obovata, these sites are located Cytogenetical studies showed than conifers possess the on chromosomes II, III, IV, V, VIII, X; in addition to them, a high amounts of DNA and large stable genomes; their minor signal has been observed in the long arm of karyotypes include large and similar on size chromosomes. chromosome IX. In P. schrenkiana, the major sites of 45S They developed in diploid level; polyploidy has played little rDNA are located in chromosomes III, IV, V, VIII, X; minor role in their evolution (Khoshoo 1959, 1961, 1962, Mehra sites were mapped in pericentromeric regions of 1968, Grant 1976, Delevoryas 1980, Muratova and Kruklis chromosomes II and IX. In P. schrenkiana there is on 1982, Kozubov and Muratova 1986, Ohri and Khoshoo 1986, chromosome II has a minor site, while P. obovata has a major Hizume 1988, Murray 1998, 2013, Ahuja 2005). Different site. species of conifers have not large differences in the average

Clusters of 5S rRNA genes occur on two arms of frequency of chiasmata per bivalent and per cell (Muratova, chromosome III of the both species; this chromosome also 2019). Members of Pinaceae and Cupressaceae families have contains major locus of 45S rDNA near the signal of 5S rDNA. a constant number and morphological types of chromosomes. Our and literature data show that the localization of 5S rDNA Working with such genomes, new information can be obtained in Picea karyotype is different from that of other Pinaceae using molecular cytogenetic markers. The use of molecular

10x90.

Comparison of two closely related Picea species – P.

obovata and P. schrenkiana, was carried out using the FISH

method.

a

b c


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Fig. 10. Comparative idiogram of Picea schrenkiana (first chromosome in each pair) and P. obovata (second chromosome). colour, location of major sites of 45S rRNA genes major are indicated in red colour, minor sites of 45S rRNA are indicated in pink

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I-XII – the numbers of the chromosomes.


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Fruitification of forest species in Siberia. Novosibirsk, Nauka,


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